Top-down control of phytoplankton in a shallow hypertrophic lake: Little Mere (England)

Top-down control of phytoplankton by zooplankton is possible through reductions in density of zooplanktivorous fish. Little Mere is a shallow lake where the effects of sewage effluent caused such a reduction. This allowed the large-bodied cladoceran, Daphnia magna Straus, to develop huge populations, preventing potentially large algal crops from developing.Subsequent diversion of the effluent is anticipated to lead to recovery of the fish community, reduced numbers of large-bodied grazers, and increased phytoplankton biomass. Whether the aquatic plant community, present in Little Mere, is resilient to such changes may depend upon whether cyanophytes are favoured, or not.     

Mesocosm experiments on the interaction of sediment influence, fish predation and aquatic plants with the structure of phytoplankton and zooplankton communities

1. Little Mere, U.K., received large quantities of sewage effluent until 1991, when the effluent was diverted. Experiments, carried out in mesocosms in 1992 and 1993, were designed to predict the effects of: (i) reduced external nutrient loading; (ii) reduced internal loading from the sediment; and (iii) recolonization by fish of the better aerated water. Treatments included isolation of the water from the underlying sediment or exposure to the sediment (which lacked plants in 1992, but was covered by Potamogeton berchtoldii in 1993) crossed with different population densities of Rutilus rutilus in 1992 and of Perca fluviatilis in 1993.
2. Exposure to sediment (as opposed to isolation from it) resulted in no net change in the biovolumes of most major algal groups, but this masked major complementary effects on individual species. The experiments showed a decreasing influence of the sediment, between 1992 and 1993, in determining water chemistry, and an increasing pH between years but no increase in cyanophyte dominance. This had been anticipated because a lake upstream provides abundant inocula, and conditions in Little Mere after diversion of effluent were expected to favour cyanophytes.
3. Roach and perch additions to the mesocosms resulted in reductions in Daphnia populations but increases in numbers of small Cladocera and copepods. Plant-associated Cladocera were unaffected by fish. The presence of submerged plants to some extent reduced fish predation effects on Daphnia hyalina .
4. The experimental results in general accurately predicted subsequent events in the open lake.     

Mesocosm experiments on the interaction of sediment influence, fish predation and aquatic plants with the structure of phytoplankton and zooplankton communities

1. Little Mere, U.K., received large quantities of sewage effluent until 1991, when the effluent was diverted. Experiments, carried out in mesocosms in 1992 and 1993, were designed to predict the effects of: (i) reduced external nutrient loading; (ii) reduced internal loading from the sediment; and (iii) recolonization by fish of the better aerated water. Treatments included isolation of the water from the underlying sediment or exposure to the sediment (which lacked plants in 1992, but was covered by Potamogeton berchtoldii in 1993) crossed with different population densities of Rutilus rutilus in 1992 and of Perca fluviatilis in 1993.
2. Exposure to sediment (as opposed to isolation from it) resulted in no net change in the biovolumes of most major algal groups, but this masked major complementary effects on individual species. The experiments showed a decreasing influence of the sediment, between 1992 and 1993, in determining water chemistry, and an increasing pH between years but no increase in cyanophyte dominance. This had been anticipated because a lake upstream provides abundant inocula, and conditions in Little Mere after diversion of effluent were expected to favour cyanophytes.
3. Roach and perch additions to the mesocosms resulted in reductions in Daphnia populations but increases in numbers of small Cladocera and copepods. Plant-associated Cladocera were unaffected by fish. The presence of submerged plants to some extent reduced fish predation effects on Daphnia hyalina .
4. The experimental results in general accurately predicted subsequent events in the open lake.     

Changes in a deep lake following sewage diversion – a challenge to the orthodoxy of external phosphorus control as a restoration strategy?

1. The restoration of deep lakes has traditionally focused on reducing the external phosphorus loading. 2. Following the diversion of sewage effluent, that led to marked reductions in nutrient concentrations in its main inflow, Rostherne Mere has shown no reduction in phosphorus or chlorophyll a concentrations. A shallow lake upstream (Little Mere), however, has shown a marked response to effluent diversion. 3. Nutrient budgets for Rostherne Mere reveal that sewage effluent was by far the most significant external source of total phosphorus and that diffuse drainage from the catchment was the most significant external source of dissolved inorganic nitrogen. Phosphorus loads from groundwater and a bird roost were insignificant. Internal sources of phosphorus were, however, considerable and were largely responsible for the observed delay in recovery. 4. Phosphorus limitation of phytoplankton biomass may never be attainable because of substantial internal and diffuse sources of phosphorus, combined with a long retention time. Nitrogen is likely to be more important in limiting phytoplankton biomass. Control of diffuse nitrogen sources may therefore be more effective in the restoration of the deeper lakes of this region.     

Two mesocosm experiments investigating the control of summer phytoplankton growth in a small shallow lake

1. Mesocosm experiments were carried out to examine the relative importance of top down (fish predation) and bottom up (nutrient addition) controls on phytoplankton abundance in a small shallow lake, Little Mere, U.K., in 1998 and 1999. These experiments were part of a series at six sites across Europe. 2. In the 1998 experiment, top‐down processes (through grazing of large Cladocera) were important in determining phytoplankton biomass. The lack of plant refugia for zooplankton was probably important in causing an increasing chlorophyll a concentration even at intermediate fish density. Little Mere normally has abundant macrophytes but they failed to develop substantially during both years. Bottom‐up control was not important in 1998, most probably because of high background nutrient concentrations, as a result of nutrient release from the sediments. 3. In 1999 neither top‐down nor bottom‐up processes were significant in determining phytoplankton biomass. Large cladoceran grazers were absent even in the fish‐free enclosures, probably because dominance of cyanobacteria and high phytoplankton biomass made feeding conditions unsuitable. As in 1998, bottom‐up control of phytoplankton was not important, owing to background nutrient concentrations that were even higher in 1999 than in 1998, perhaps because of the warmer, sunnier weather. 4. The differing outcomes of the two experiments in the same lake with similar experimental designs highlight the importance of starting conditions. These conditions in turn depended on overall weather conditions prior to the experiments.     

Consequences of reduced nutrient loading on a lake system in a lowland catchment: deviations from the norm?

1. Lake restoration from eutrophication often rests on a simple paradigm that restriction of phosphorus sources will result in recovery of former relatively clear‐water states. This view has apparently arisen from early successful restorations of deep lakes in catchments of poorly weathered rocks. Lakes in the lowlands, however, particularly shallow ones, have proved less tractable to restoration. This study of three lowland lakes provides insights that illuminate a more complex picture. 2. The lakes lie in a sequence along a single stream in a mixed urban and rural landscape. Severely deoxygenating effluent from an overloaded sewage treatment works was diverted from the catchment in 1991. Effects on two lakes, Little Mere (z_max <2 m) and Rostherne Mere (z_max 31 m) were followed until 2002. Mere Mere (z_max = 8 m), upstream of the former works, acted as a comparison for changes in water chemistry. Mere Mere showed no change in total phosphorus (TP), total inorganic nitrogen, or planktonic chlorophyll a concentrations. Increased winter rainfall was associated with higher winter soluble reactive phosphorus (SRP) and ammonium concentrations in its water. 3. Little Mere changed from a deoxygenated, highly enriched, fishless system, with large populations of Daphnia magna Straus, clear water and about 40% aquatic plant cover, to a slightly less clear system following diversion. Daphnia magna was replaced by D. hyalina Leydig as fish recolonised. Spring peaks of chlorophyll a declined but summer concentrations increased significantly. Annual mean chlorophyll a concentrations thus showed no change. Submerged plants became more abundant (up to 100% cover), with fluctuating community composition from year to year. Summer release of SRP from the sediment was substantial and has not decreased since 1993. The summer phytoplankton was apparently controlled by nitrogen availability perhaps with some influence of zooplankton grazing. SRP was always very abundant. The lake appeared to have reached a quasi‐stable state by 2002. 5. Rostherne Mere showed a steady decline in TP and SRP concentrations following effluent diversion apparently as a result of steady dilution by water with lower phosphorus concentration. Decline in phosphorus concentrations was much less rapid than expected because of internal remobilisation from the hypolimnion and sediments. There have been no changes in chlorophyll a concentration or of nitrogen availability and by 2002 the phytoplankton probably remained limited by a combination of mixing, grazing and nitrogen. 6. A seeming paradox is, thus, that immense changes in phosphorus budgets have shown no consequences for phytoplankton chlorophyll concentrations in either of the lakes, although the seasonal distribution has altered in Little Mere. Although these case studies deviate from others, for both shallow and deep lakes, they represent distinctive situations rather than undermining conventional models.     

Response of a shallow Mediterranean lake to nutrient diversion: does it follow similar patterns as in northern shallow lakes?

1. In view of the paucity of data on the response of warm shallow lakes to reductions in nutrient loading, this paper presents a long‐term limnological data set to document changes in the food‐web of a shallow Mediterranean lake (Lake Albufera, Valencia, Spain) that has experienced reductions in phosphorus (P) (77%) and nitrogen (N) (24%) loading following sewage diversion. 2. Nine years after sewage diversion, P concentration in the lake was reduced by 30% but remained high (TP = 0.34 mg L^?1), although the mean water retention time in the lake was only 0.1 years. Nitrate concentrations did not significantly change, probably because the lake continued to receive untreated effluents from ricefields. 3. Chlorophyll a concentration was reduced by half (annual mean of 180 μg L^?1). Cyanobacteria abundance remained high but its composition changed towards smaller species, both filamentous and chroococcal forms. 4. Cladocera abundance increased and reached peaks twice a year (December to March and July to September). After nutrient reduction, short‐term clear‐water phases (up to 5 weeks) occurred during February to March in several years, concomitant with annual flushing of the lake and lower fish densities. The abundance of Cladocera in winter contrasted with the spring peaks observed in northern restored shallow lakes. The zooplankton to phytoplankton biomass ratio remained lower than in northern temperate shallow lakes, probably because of fish predation on zooplankton. 5. Improvement of the water quality of Lake Albufera remained insufficient to counteract littoral reed regression or improve underwater light allowing submerged plants re‐colonise the lake. 6. Sewage diversion from Lake Albufera impacted the food web through the plankton, but higher trophic levels, such as fish and waterfowl, were affected to a lesser degree. Although the fish species present in the lake are mainly omnivorous, long‐term data on commercial fish captures indicated that fish communities changed in response to nutrient level and trophic structure as has been observed in restored shallow lakes at northern latitudes. 7. Phosphorus concentrations produced similar phytoplankton biomass in Lake Albufera as in more northern shallow lakes with abundant planktivorous fish and small zooplankton. However, in Lake Albufera, high average concentrations were maintained throughout the year. Overall, results suggest that nutrient control may be a greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes.     

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Short-term and long-term effects of zooplanktivorous fish removal in a shallow lake: a synthesis of 15 years of data from Lake Zwemlust

SUMMARY 1. Removal of zooplanktivorous fish (mainly bream) in 1987 from a shallow eutrophic lake in the Netherlands, Lake Zwemlust, resulted in a quick switch from a turbid state with cyanobacteria blooms to a clear state dominated by macrophytes.
2. The clear state was not stable in the long term, however, because of high nutrient loadings.
3. In 1999, another removal of zooplanktivorous fish (mainly rudd) had similar effects as in 1987, although macrophytes returned more slowly.
4. In the years directly following both interventions there was a 'transition period' of very clear water with high densities of zooplanktonic grazers in the absence of macrophytes; low oxygen concentrations indicate that during those years primary production was low relative to heterotrophic activity.
5. The transition period appears to provide the light climate necessary for the return of macrophytes.
6. Reduction of nutrient loading is necessary to improve water quality in Lake Zwemlust in the long term. In the short term, repeated fish stock reduction is a reasonable management strategy to keep Lake Zwemlust clear.     

The effects of tench (Tinca tinca (L.)) and sticklebacks (Gasterosteus aculeatus L.) on planktonic and benthic communities in mesocosms in a shallow lake

The effects of introducing a zooplanktivorous fish, three-spined stickleback, (Gasterosteus aculeatus) and a benthivorous fish, tench (Tinca tinca) separately and in combination to replicated experimental enclosures with two density levels of white water lily (Nymphaea alba) were studied in Little Mere, UK. Numbers of Daphnia hyalina were high and only slightly diminished at reduced lily densities, probably due to stickleback predation, but there was no consequential effect on phytoplanktonic chlorophyll a concentrations. Tench reduced the numbers of gastropods but not of other macroinvertebrates, and in turn increased the biomass of periphyton growing on artificial substrata within the enclosures. The higher lily density reduced oxygen concentrations and pH values and increased total phosphorus and soluble reactive phosphorus concentrations but otherwise had little effect on water chemistry. There was little interactive effect of the fish species. The results are integrated with those of six other such enclosure experiments carried out in Little Mere since 1992.     

Habitat architecture and trophic interaction strength in a river: riffle-scale effects

Invertebrate algal grazer densities were manipulated in a temperate river to discover the impact of differences in riffle-scale architectural complexity on the strength of the trophic interaction between grazers and epilithic algae. Animal densities were manipulated by manual removal in architecturally complex boulder-cobble riffles and simpler bedrock riffles, with the complexity of smaller-scale architecture held constant. Responses in algal density were recorded before and after a month of manipulations, together with grazer colonization rate and body sizes. The experiment was carried out in winter and again in summer. The interaction between grazers and algae differed between habitats and seasons. In winter, when algae were growing, the more complex rifflescale architecture in the boulder-cobble riffles created a refuge from grazing for algae. This was probably the result of the movement abilities of the grazers interacting with habitat architecture, and potentially also due to the control of predatory fish densities by habitat architecture resulting in greater predation pressure on grazers in boulder-cobble riffles. Therefore the impact of highly complex riffle-scale architecture was to weaken the strength of the trophic interaction between algae and their grazers by reducing grazer densities, while potentially strengthening the trophic interaction between grazers and their fish predators.     

The relative importance of top-down and bottom-up control of phytoplankton in a shallow macrophyte-dominated lake

1. Mechanisms stabilizing the plant-dominated clear-water state were investigated in Little Mere, U.K. Replicated, factorial, mesocosm experiments, carried out in 1995 and 1996, were designed to investigate the relative importance of top-down (zooplankton grazing) and bottom-up (nitrogen-limitation) control in limiting algal growth, and the role of macrophytes in these processes. Treatments included increased salinity (1995) and sticklebacks (1996) to reduce zooplankton numbers, weekly nitrate additions and removal of macrophytes. 2. Contrary to the results of other studies, submerged plants did not reduce nitrate concentrations. Owing to the high stickleback density in the enclosures with fish, macrophytes did not provide a refuge for zooplankton during the experiment. In Little Mere, however, where fish densities are lower, macrophytes probably play a key role in maintaining clear water by providing refuge for pelagic zooplankton and habitat for attached Cladocera. 3. Phytoplankton in Little Mere was not nitrogen- (N) limited during the growing season. Although nitrogen availability sets a maximum potential phytoplankton biomass it was not realized owing to control by zooplankton grazing.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

Consumer-dependent responses of lake ecosystems to nutrient loading

The nutrient loading concept proposes that algal biomass, waterclarity and the processes of lake eutrophication are a functionof nutrient loading. We hypothesized that grazers play an importantrole in determining the impacts of nutrient loading on algalbiomass and water clarity, and the overall eutrophication process.To test how the contrasting grazer communities modify the fateof nutrients, we added nutrients (nitrate and phosphate) ata known loading rate to four large enclosures, but in two ofthe four enclosures large cladoceran grazers (Daphnia >1mm mean length) were allowed to develop by removing the planktivorousfish. In the remaining two enclosures, the development of largeDaphnia was prevented by adding planktivorous fish. The concentrationsof epilimnetic total phosphorus (TP) increased at a similarrate in all four enclosures. However, the daily accumulationof added phosphate into the participate or planktonic forms,especially into plankton <20 µm, was three times fasterwhen large Daphnia were absent than when large Daphnia wereabundant. In the enclosures with large Daphnia, added phosphatewas accumulated in the dissolved pool instead. At a constantnutrient loading, algal biomass (chlorophyll a) increased fourtimes faster in the enclosures without large Daphnia than inthose with large Daphnia. Similarly, Secchi depth declined from3.5 to <1 m when Daphnia were absent, but did not declinewhen Daphnia were common. Our results demonstrate that the samenutrient loading and the resultant increase in epilimnetic TPdo not produce the same trophic conditions, as indicated byalgal biomass and water clarity, if the grazers of the majorassimilators of nutrients (the fraction of plankton edible toDaphnia) are different. We suggest that stratified lake ecosystemshaving functionally dominant large grazer communities may beless prone to eutrophication than those lacking large grazers.Consistent with the nutrient loading concept, epilimnetic concentrationsof phosphorus increase proportionately with increased loadingof phosphorus, but the trophic conditions of ecosystems indicatedby algal biomass and water clarity do not follow the same patternsunder contrasting conditions of grazer communities. We suggestthat models predicting algal biomass from loading rates shouldaccount for the role of grazers.     

Cascading effects of predatory fish on the composition of benthic algae in high‐altitude streams

Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.     

The Effects of Stocking and Removal of a Nonnative Salmonid on the Plankton of an Alpine Lake

Bighorn Lake, a fishless alpine lake, was stocked with nonnative brook trout, Salvelinus fontinalis, in 1965 and 1966. The newly introduced trout rapidly eliminated the large crustaceans Hesperodiaptomus arcticus and Daphnia middendorffiana from the plankton. In July 1997, we began to remove the fish using gill nets. The population comprised 261 fish that averaged 214 g in wet weight and 273 mm in fork length. Thereafter, zooplankton abundance increased within weeks. Early increases were caused by the maturation of Diacyclops bicuspidatus, few of which reached copepodid stages before the removal of the fish because of fish predation. Daphnia middendorffiana, absent when fish were present, reappeared in 1998. Hesperodiaptomus arcticus, which had been eliminated by the stocked fish, did not return. The proportion of large zooplankton increased after fish removal, but their overall biomass did not change. Algal biomass was low and variable throughout the 1990s and correlated with water temperature but not with nutrient concentrations or grazer densities. Diatoms were the most abundant algal taxon in the lake, followed by Dinophyceae. Chrysophyceans and cryptophyceans were eliminated after the fish were removed. Chlorophyll a concentrations were unaffected. Gill netting is a viable fish eradication technique for smaller (less than 10 ha), shallow (less than 10 m deep) lakes that lack habitable inflows and outflows or other sensitive species. Further work is required to define appropriate removal methods for larger lakes and watersheds. Received 30 May 2000; Accepted 14 November 2000.     

Restoration of the eutrophic Lake Eymir, Turkey, by biomanipulation after a major external nutrient control I

Nutrient loading in lakes is recognized as a serious threat to water quality. Over 25 years of raw sewage effluent discharge shifted Lake Eymir from a state dominated by submerged plants to a turbid water state. Successful effluent diversion undertaken in 1995 achieved 88% and 95% reductions in the areal loading of total phosphorus (TP) and dissolved inorganic nitrogen (DIN), respectively. Furthermore, the reduced load of TP was very close to the suggested threshold areal load (0.6 g m^–2 yr^–1) to attain recovery. Even though diversion also reduced the in-lake TP level by half, the poor water clarity and low submerged plant coverage (112 ± 43 cm and 2.5% coverage of the lake total surface area, respectively) persisted. Domination of the fish stock by planktivorous tench (Tinca tinca L.) and the benthivorous common carp (Cyprinus carpio L.) (66 ± 0.7 and 31 ± 1 kg CPUE, respectively) appeared to perpetuate the poor water condition. A substantial fish removal effort over 1 year achieved a 57% reduction in the fish stock which led to a 2.5-fold increase in Secchi disk transparency. This increase occurred largely because of a 4.5-fold decrease in the inorganic suspended solid concentration, and to some extent, a decrease in chlorophyll-a concentration. A strong top-down effect of fish on the large-sized grazers was evident as density and the body size of Daphnia pulexde Geer increased significantly after the fish removal. Even though the spring and annual euphotic depths occurred well above the maximum and mean depths of the lake, respectively, re-development of submerged plants was poor (6.2% coverage). A weak re-establishment of submerged plants might be attributed to an insufficiently viable seed bank, inappropriate chemical conditions of the sediment (severe oxygen deficiency), or to the high coot (Fulica atra L.) density. However, the top-down effect of fish appeared to be of great importance in determining water clarity, and in turn, conditions for submerged plant development in a warm temperate lake as recorded in the north temperate lakes. Furthermore, this study provides evidence for the importance of top-down control of fish, which, in turn, can be effectively utilised as a restoration strategy in warm-temperate lakes as well. More applications, along with long monitoring programs, are needed to develop a better understanding about requirements for biomanipulation success in this climate.     

Periphyton-macroinvertebrate interactions in light and fish manipulated enclosures in a clear and a turbid shallow lake

In a clear and a turbid freshwater lake the biomasses of phytoplankton, periphytic algae and periphytonassociated macrograzers were followed in enclosures with and without fish (Rutilus rutilus) and four light levels (100%, 55%, 7% and < 1% of incoming light), respectively. Fish and light affected the biomass of primary producers and the benthic grazers in both lakes. The biomass of primary producers was generally higher in the turbid than the clear lake, and in both lakes fish positively affected the biomass, while shading reduced it. Total biomass of benthic grazing invertebrates was higher in the clear than in the turbid lake and the lakes were dominated by snails and chironomids + ostracods, respectively. While light had no effect on the biomass of grazers in the clear lake, snail breeding was delayed in the most shaded enclosures and presence of fish reduced the number of snails and the total biomass of grazers. In the turbid lake ostracod abundance was not influenced by light, but was higher in fish-free enclosures. Density of chironomids correlated positively with periphyton biomass in summer, while fish had no effect. Generally, light-mediated regulation of primary producers was stronger in the turbid than in the clear lake, but the regulation did not nambiguously influence the primary consumers. However, regulation by fish of the benthic grazer community was stronger in the clear than in the turbid lake, and in both lakes strong top-down effects on periphyton were seen. The results indicate that if present-day climate in Denmark in the future is found in coastal areas at higher latitudes, the effect of lower light during winter in such areas will be highest in clear lakes, with typically lower fish biomass and higher invertebrate grazer density.     

Bottom-up and cascading top-down control of macroalgae along a depth gradient

On marine rocky shores, macroalgal herbivory is often intense, such that the cascading effects of fish predation may contribute to the control of algal communities. To estimate the magnitudes of top-down and bottom-up control on a macroalgal community, we manipulated the access of carnivorous fish to macroalgal colonization substrates, as well as nutrient availability, at two sub-littoral depths. There were three levels of fish manipulation: natural fish community, no fish and the enclosure of one common species, the perch, Perca fluviatilis. We found a clear cascade effect of fish predation on both the total density and several individual species of macroalgae, which was more pronounced in deep than shallow water. The density of the dominant grazers, i.e. snails, increased in nutrient-enriched conditions; perch were inefficient in controlling herbivores, and had therefore no cascading effect on algal densities under such conditions. Although nutrients enhanced the growth of opportunistic algae, herbivores, in the absence of fish, inhibited this response. While algal diversity was higher in shallow than in deep water, the enrichment effect was opposite at the two depths with lowered diversity in the shallows and increased at depth. Our results indicate that fish predation is an efficient regulator of meso-herbivores and that its effect thereby cascades onto the producer trophic level such that both perennial and opportunistic algae benefit from the presence of fish. This cascade effect is probably stronger at depth where predation efficiency is less disturbed by wave motion.